Clearwing Moths of Australia and New Zealand
(Lepidoptera: Sesiidae)
m
W. DONALD DUCKWORTH and
THOMAS D. EICHLIN
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 180
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y • N U M B E R 1 8 0
Clearwing Moths of Australia and New Zealand
(Lepidoptera: Sesiidae)
W. Donald Duckworth and Thomas D. Eichlin
SMITHSONIAN INSTITUTION PRESS City of Washington
1974
ABSTRACT
Duckworth, W. Donald, and Thomas D. Eichlin. Clearwing Moths of Australia and New Zealand (Lepidoptera: Sesiidae). Smithsonian Contributions to Zool- ogy, number 180, 45 pages, 50 figures, 6 maps, 1974.—The family Sesiidae in Australia and New Zealand is revised and one new species is described. The history, biology, geographical distribution, morphology, and classification of the family are discussed in detail. All the species are reviewed regarding their taxo- nomic history, distribution, identity, and morphology. A key to species and higher categories is provided. Distribution maps, photographs of adults, illustra- tions of the male and female genitalia, wing venation, and other aspects of morphology are included.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5129. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data Duckworth, W. Donald
Clearwing Moths of Australia and New Zealand (Lepidoptera: Sesiidae) (Smithsonian contributions to zoology, no. 180)
Supt. of Docs, no.: SI 1.27: 180.
1. Aegeriidae. 2. Insects—Australia. 3. Insects—New Zealand. I. Eichlin, Thomas D., 1938- joint author. II. Title. HI. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 180.
QL1.S54 no. 180 [QL561.A34] 591'.08s [595.7'81] 74-6058
For sale by the Superintendent of Document!), U.S. Government Printing Office Washington, D.C. 20402 - Price $1.15 (paper cover)
Contents
Page
Introduction \ History 2 Morphology 3 Biology 5 Geographical Distribution 6 Classification 8 Taxa Transferred to Other Families 9 Family Stathmopodidae 9 Genus Dolophrosyne Durrant 9 Dolophrosyne balteata Durrant 9 Family Glyphipterygidae 9 Genus Sagalassa Walker 9
Sagalassa homotona (Swinhoe)7 new combination 9 Checklist 9 Family Sesiidae 10 Key to the Species and Higher Categories 11 Subfamily Tinthiinae LeCerf 12 Tribe Pennisetiini Naumann 12 Genus Pennisetia Dehne 12 Pennisetia igniflua (Lucas), new combination 12 Pennisetia eusphyra (Turner), new combination 14 Tribe Tinthiini LeCerf 15 Genus Tinthia Walker 15 Tinthia xanthospila Hampson 15 Subfamily Paranthreninae Niculescu 16 Tribe Paranthrenini Niculescu 16 Genus Albuna Edwards 16 Albuna carulifera (Hampson), new combination 16 Albuna isozona (Meyrick), new combination 18 Albuna oberthuri (LeCerf), new combination 19 Albuna zoniota (Turner), new combination 21 Subfamily Sesiinae Boisduval 21 Tribe Melittini LeCerf 22 Genus Melittia Hubner 22 Melittia amboinensis Felder 22 Melittia chalybescens Miskin 24 Tribe Synanthedonini Niculescu . . 25 Genus Synanthedon Hubner 25 Synanthedon tipuliformis (Clerck) . 25 Synanthedon cupreifascia (Miskin) 26 Genus Carmenta Edwards 28 Carmenta chrysophanes (Meyrick), new combination 28 Carmenta commoni, new species . . 30 Carmenta xanthogyna (Hampson), new combination 31 Literature Cited 32
iii
Clearwing Moths of Australia and New Zealand
(Lepidoptera: Sesiidae)
W. Donald Duckworth and Thomas D. Eichlin
Introduction
The Sesiidae ( = Aegeriidae) is a well-defined family, worldwide in distribution. Consisting of over 1000 described species, the adult members of this family are primarily diurnal and highly mi- metic in both appearance and behavior. The de- gree to which species in this family have become modified structurally and behaviorally to resemble their models, primarily bees and wasps, is truly remarkable. The wings are narrow and, in most instances, partially devoid of scales; the abdomen is commonly banded with orange or yellow and narrowed basally either by actual constriction or scaled so as to create an illusion of constriction.
In addition, the legs are usually modified to re- semble those of the model even to the extent, in some groups, of scale tufts tipped in yellow to simulate the pollen-gathering devices of certain bees.
The larvae are primarily borers in the trunks, bark, stems, or roots of trees, shrubs, and vines or in the stems and roots of herbaceous plants. Some species are inquiline borers in galls on woody and herbaceous plants. Although a great deal of bio- logical data is known for species that occur in tem-
W. Donald Duckworth, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Thomas D. Eichlin, Laboratory Services/
Entomology, Division of Plant Industry, California Depart- ment of Food and Agriculture, Sacramento, California 958N.
perate regions, particularly the Nearctic and Pale- arctic, the life histories of species in tropical and subtropical areas are still very incompletely known.
The present study is an outgrowth of research currently underway by the authors on the Western Hemisphere sesiids and represents the first detailed account of all the species known to occur in Aus- tralia and New Zealand. As presently defined, the sesiid fauna of this area consists of 14 species in 6 genera, of which one species is described as new, and one, Synanthedon tipuliformis (Clerck), is an introduced species from Europe. In addition, two species, Balataea homotona Swinhoe and Doloph- rosyne balteata Durrant, previously assigned to the family have been found to be improperly placed and are herein transferred to other families.
Throughout the course of this study we have been handicapped by the small number of speci- mens available for study. This is a fairly general situation for the sesiids throughout the world and is undoubtedly due, in large part, to the diurnal flight period and fugitive behavior of the adults, coupled with the endophagous boring habit of the larvae. As a result, most collections are sorely lacking in sufficient material to adequately support modern systematic studies. It is our hope that this paper will serve to stimulate Australian biologists to focus more attention on this unique family of moths.
ACKNOWLEDGMENTS.—The authors wish to ac- knowledge with gratitude the cooperation and assist- 1
ance of the following individuals and institutions that, through their support and encouragement, have materially aided the present study: Dr. I. F. B.
Common, Commonwealth Scientific and Industrial Research Organization, Canberra, Australia; Dr.
Gordon F. Gross, South Australian Museum, Ade- laide; Mr. A. Neboiss, National Museum of Vic- toria, Melbourne, Australia; Dr. C. N. Smithers, Australian Museum, Sydney; Mr. J. S. Dugdale, Department of Scientific and Industrial Research, Nelson, New Zealand; Mr. Paul E. S. Whalley, Dr.
Klaus Sattler, British Museum (Natural History);
Mr. I. Lansbury, Hope Department of Zoology, University of Oxford; Dr. P. Viette, Museum Na- tional D'Histoire Naturelle, Paris, for lending for study types and other specimens in their charge.
We wish to particularly acknowledge our grati- tude to Dr. Ian F. B. Common, CSIRO, whose will- ingness to share with us his extensive knowledge of Australian Lepidoptera, both through prompt response to our numerous inquiries by mail and directly during a brief visit by one of the authors (Duckworth) to Australia, significantly contrib- uted to the completion of the project.
The authors also wish to acknowledge the assist- ance of Mr. George Venable, departmental illus- trator, for the illustrations and maps; Mrs. Vera Milbank, technician, for bibliographic assistance;
Mr. Timothy P. Friedlander, undergraduate fellow, for assistance in all phases of the study; and the National Museum of Natural History Photo- graphic Laboratory for the photographs.
This study was aided in part by a grant from the Secretary's Fluid Research Fund, Smithsonian Re- search Foundation number 112-715, and a Travel Grant from The Entomological Society of America.
History
As one might expect, considering the limited number of species and infrequency of capture, the previous literature concerning the Australian and New Zealand sesiids is anything but profuse. The initial documentation of the family for this region was by Meyrick (1886) and consisted of the descrip- tion of two new species (Sesia isozona and Sesia chrysophanes) from Australia, and Synanthedon tipuliformis (Clerck) listed as an introduced species in New Zealand.
Turner (1917), in a paper describing a number of new Australian species in various families, pre- sented a summary of the family Aegeriidae (= Sesiidae) noting that "the known species are of Indo-Malayan origin." Five years later Turner (1922) summarized the family once again in order to include new taxa from Australia resulting from the publication of intervening papers by LeCerf (1917) and Hampson (1919). Although both of the latter papers were primarily concerned with other faunal areas, both included species from Aus- tralia and, even more important, higher category concepts of significance to the Australian region.
With the publication of Turner's paper, the known sesiid fauna consisted of 15 species in 7 genera, all (except the introduced S. tipuliformis) from Queensland and northern Australia.
In contrast to Turner's figures, Dalla Torre and Strand (1925) list 17 species in 11 genera as occur- ring in Australia, and Gaede (1933) lists 14 species in 8 genera. These publications were worldwide treatments of the family and illustrate the incon- sistencies inherent in such ambitious undertakings.
Attempts at family definition for the Australian sesiids were provided by Tillyard (1926) and Common (1970). In both instances only a family level treatment was intended and no more than a few selected species were included.
In addition to the works previously cited, it is important to note a number of publications which, although they have not necessarily included data directly pertaining to the Australian fauna, are of considerable significance in the development of our overall knowledge of the family. Beutenmuller (1901) provided the first comprehensive review of the family in North America, including a sys- tematic review of the literature. In addition, an attempt was made to present all the known biologi- cal information concerning the included species.
This tradition was continued and expanded by Engelhardt (1946), whose later revision of the family for North America greatly expanded our knowledge of the life histories and habits of the group.
More recently, the publications of Niculescu (1964) and Naumann (1971) have contributed significantly to the higher category concepts with- in the family and have aided in the clarification of several nomenclatural problems of long standing.
Little information has been published regarding
NUMBER 180
the systematics of the immature stages of sesiids, thus MacKay's (1968) revision of the North Amer- ican species, based on late instar larvae, provides a vital resource for comparison of higher category concepts as well as for elucidation of larval morphology and species identification.
Morphology
It is not our intention to deal in great detail with the comparative morphology of the Sesiidae in the present paper. The limited nature of the Australian-New Zealand sesiid fauna, together with the lack of significant morphological diver- gence from patterns observed for the family in other parts of the world, makes it more desirable to present a detailed analysis of character trends as a part of our studies on the Western Hemi- sphere portion of the family. The family descrip- tion included in the present study provides a complete summation of structural characteristics for the group and is based on an extensive world- wide survey of the family.
In general, the sesiids are a remarkably homo- geneous group morphologically. They are readily recognized superficially by their narrow and more or less transparent wings, clavate or filiform anten- nae, and the generally elongate, slender abdomen, which is tapered posteriorly and terminated by a more or less prominent anal tuft of scales. The mimetic nature of the group is generally apparent by the pattern and scaling of the wings, by the ab- domen being frequently banded or striped with yellow or orange, and by the legs being usually long and variously tufted with scales.
While family recognition is relatively easy, char- acterization of taxa below the family level requires careful examination of various character systems (e.g., wing venation, genitalia, legs, head, etc.), for the distinguishing differences are frequently quite subtle. The keys provided in the present study are designed to facilitate identification through utili- zation of easily observed characters; however, they have been correlated with additional characters derived from intensive study of adult morphology as well as data derived from other life stages and biologies, whenever possible.
One portion of sesiid morphology displays unique modifications worthy of particular empha-
sis. The wings of sesiids are, as mentioned previ- ously, very narrow. It is presumed this reduction has resulted from selection favoring hymenopterous mimicry; however, as the wings become more nar- row the mechanisms found in most Lepidoptera for coupling the fore and hindwing during flight become less efficient, especially for the typically rapid flight exhibited by most sesiids. Thus, in the family a number of modifications in the wing- coupling structures have occurred that serve to in- sure synchronous action of the fore and hindwings.
In most moths the mechanism for wing-coupling consists of a single-spined frenulum arising from the costal sclerite at the base of the hindwing in males and a several-spined (usually three) frenu- lum in the females. The frenulum is secured to the underside of the forewing by varying devices, gen- erally termed the retinaculum. In the Sesiidae both males and females have developed a single- spined frenulum (two-spined in some females), which is secured by an overfold of subcosta that projects to near the wing membrane. Also, there is a row of scales from the radius, directed anteri- orly, which aid in holding the frenulum more firmly in place. The development of a single-spined frenulum and a subcostal retinaculum is generally considered an advanced specialization in moths and one which is generally confined to the males (Braun, 1924). The attainment of this condition by both sexes in sesiids suggests, at least in this one character, a highly derived condition. To our knowledge, no other family of moths has achieved this degree of specialization in this character thoughout its component taxa.
In addition to the advanced frenulum- retinaculum device, the sesiids have further strengthened wing-coupling through the inter- locking of the dorsal edge of the forewing with the costal edge of the hindwing. This is accomplished by the rolling downward of the dorsal margin of the forewing and locking firmly into nearly the entire length of the costal margin of the hindwing, which is similarly rolled upward. Each of these margins bears a series of interlocking, recurved spines, which further serve to secure the attach- ment.
The firm bond attained through the two methods described above has provided the sesiids with the most effective wing-coupling mechanism found in Lepidoptera and undoubledly enhances the swift
and evasive flight behavior typically observed in the family. In addition, the ability to fly rapidly would appear to be a necessary trait if hymenop- terous mimicry is to be extended to flight behavior.
That this transition has been accomplished by many species of sesiids is readily demonstrated by the numbers of specimens inadvertently captured by collectors of Hymenoptera.
It should be noted in connection with the wing- margin coupling mechanism that it is not possible at this time to adequately evaluate the potential phylogenetic implications of this development in sesiids. Braun (1919), in her study of the wing structure of Lepidoptera, comments as follows:
"The possession of the row of costal spines by many of the Trichoptera and by more primitive Lepidoptera indicates that it is a persistent primi- tive character, and as such indicates common de- scent of the two groups. The preservation of these spines in many frenate Lepidoptera is one of the few connecting links between them and the Mic- ropterygidae." This would suggest the possibility that the recurved spines which function in the wing-margin coupling mechanism in sesiids are derived from the costal spines, at least in part, and as such represent an evolutionary line in wing- coupling mechanisms based on a persistent ances- tral character. However, Braun goes on to say:
"The series of costal hooks—the hamuli—which have developed in the more specialized families of Trichoptera as a means of locking the fore and hindwing together, is a specialization not found in the Lepidoptera." Thus, if the recurved wing spines in sesiids are viewed as homologous with hamuli in Trichoptera, a view apparently held by Forbes (1923), then their presence would appear to be most likely due to parallelism. Additional studies of the development of wing-coupling mech- anisms in the Lepidoptera are needed to resolve questions of this nature.
The reduction in wing width in sesiids, espe- cially in the forewing, and the interlocking fore and hindwing margins have influenced the vena- tion patterns of the wings in several ways. There is a great reduction in the anal area of the forewing and an unusual lack of reduction in the anal area of the hindwing. In the forewing there is a reduc- tion in the number of veins in many species and the anal area is so reduced that, of the anal veins,
only 2A remains and it is frequently reduced. By contrast, the anal area of the hindwing is expan- sive with patterns that include four anal veins, an extremely significant factor phylogenetically in that the presence of 4A is considered an ancestral condition lost in most of the Lepidoptera. In ad- dition, the anal area of the hindwing, in our opinion, reflects in its venational patterns the three major lines of development in the family and provides characters of value in distinguishing the three subfamilies. The Tinthiinae appear ancestral in retaining 1A (lost and represented by a wing fold in most derived groups of Lepidoptera) but have the other three anals coalescent with 2A.
In both the Paranthreninae and Sesiinae, 1A is ab- sent and represented only by a line of scales on the wing fold. In Paranthreninae, 2A and 3A are coalescent, except at the base, and 4A is present.
The Sesiinae have 2A, a portion of 3A, and 4A is present.
There is also a unique contrast in the develop- ment of the media vein in the fore and hindwing of sesiids. In the forewing the stem of media is lost, as it is in most groups of advanced Lepidoptera.
In the hindwing, however, the stem of media is present and has shifted in position toward the costal margin. The three veins normally promi- nent along the costal margin (Sc, R^ R.) are greatly reduced, closely parallel, and concealed within the upfolded costal edge. This condition led early workers to erroneously conclude that these veins were missing in the hindwings of sesiids.
The venation patterns in the wings of sesiids have proved to be of value in delimiting taxa at virtually every level within the family and are de- tailed in the descriptions of these taxa in the taxonomic portion of this study. However, the homogeneity alluded to earlier in this discussion prevails also in wing morphology and was perhaps best summarized by Comstock (in Beutenmuller, 1901): "Although the members of this family in the course of their evolution have progressed far from the primitive type of the order, they have kept closely together; there is much less variation in the structure of the wings than one would expect to find in a group so highly specialized." Based on our studies of the family to this point, much the same may be said for most character systems found in sesiids.
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Biology
Host-plant associations and other biological data are largely lacking for Australian sesiids. However, our current knowledge of seemingly general habits within the family, coupled with biological data accumulated for species in other parts of the world belonging to genera which occur in Australia, is such that certain patterns may be noted and some extrapolation made. Specific information for each Australian species, when known, is included in the descriptive portion of this study.
ADULT.—The most obvious and significant char- acteristic of clearwing moths is their structural and behavioral modifications related to mimetic asso- ciations. While this obvious adaptation is readily recognized by researchers, both past and present, virtually no investigations of a serious nature have been conducted on specific mimic-model relation- ships within the family. Thus, it is not possible to detail the biological implications of clearwing moth mimicry in other than speculative terms.
This area of sesiid biology offers numerous oppor- tunities for productive research on mimicry, adaptive aspects of behavior, and other related phenomena of broad biological significance and, hopefully, will become more widely investigated as the taxonomy of the family is clarified.
In general, clearwing moths are diurnal in adult activity periods, possess behavioral patterns asso- ciated with their models, primarily bees and wasps, and do not appear to disperse far from their host- plants. Previous studies on the behavior of selected species in North America (Girault, 1907; Gossard and King, 1918; Nielsen and Balderston, 1973) indicate that chemical sex attractants are utilized by sesiids for mate location. In addition, Nielsen and Balderston (1973) reported evidence for inter- generic sex attraction among sesiids, suggesting that more than one species may utilize the same pheromone(s) for locating mates. This finding, in turn, suggests that additional behavioral, visual, or chemical stimuli, which selects against noncon- specific mates, must come into play once physical proximity is achieved.
Once mating is accomplished, oviposition begins immediately, the eggs generally being deposited singly on or near the host-plant. The eggs are ovate, disc-shaped with flat bottoms, brown in color, with the surfaces weakly sculptured with
minute shagreening in hexagonal designs. The time required for incubation varies considerably between species and is probably dependent upon temperature. Upon hatching, the larvae bore into the host-plant directly or after migrating to another area of the plant.
LARVA.—Insofar as is known all sesiid larvae are true borers and, consequently, exhibit the lack of body color and pattern characteristic of lepidopter- ous larvae with this habit. The time required for development and the number of instars varies con- siderably, with three years and seven instars being the maximum period and number recorded for a given species. Many species in North America are univoltine and overwinter as late instar larvae, either in the plant or in chambers in the soil.
Host-plant specificity also varies considerably with- in the family. Some species and/or genera exhibit rather narrow host preferences, while others appear to exploit a broad range of plants. While available data are sketchy, Australian sesiids appear to fol- low the same pattern. For example, the genus Melittia Hubner occurs worldwide and is, so far as is known, host specific on plants in the family Cucurbi- taceae. Although the two Australian species, Melit- tiachalybescens Miskin and Melittia amboinensis Felder, have not been reared in Australia as yet, the latter species has been reported from cucurbi- taceous plants in India (David and Subramaniam, 1962), and it seems reasonable to assume the same association will be found to occur in Australia when appropriate rearings are accomplished. By contrast, Carmenta chrysophanes is capable of utilizing a number of different host-plants, which may explain, at least in part, the fact that it is the only native species of the family that is distributed into the more temperate area of Australia.
PUPA.—Pupation occurs in most species of Sesx- idae in the larval gallery. Prior to pupation the larva generally tunnels to the surface, leaving only a thin exterior covering, then retreats some dis- tance and prepares a pupal case of bits of plant material and frass. At the time of emergence the pupa moves up the gallery, penetrates the thin covering, extrudes up to two-thirds its length from the gallery, and in this position the adult emerges.
This process is facilitated by the retention of free- dom of movement in abdominal segments 3 to 7 in the male pupa and 3 to 6 in the female pupa, and by the presence of special structures on the head
and abdomen. The head is heavily sclerotized ante- riorly and usually has many ridges and projections.
The clypeus often bears a chisel-like projection that is utilized to penetrate the exit region. Move- ment through the gallery and anchoring during emergence are aided by transverse rows of dorsal spines on abdominal segments 2 to 7 in the males and 2 to 6 in the females, the females having a sin- gle row on segment 7 and both sexes having single rows on segments 8, 9, and 10. There are also large spines ventrad on segment 10 and the cremaster is lacking.
NATURAL ENEMIES.—Records of parasites of sesiids are not abundant and those which have been documented are generally in relation to species of economic concern to man. To date, there are no records concerning parasites, predators, or diseases of Australian sesiids. From other areas of the world, however, data indicate parasites in the orders Hymenoptera and Diptera; birds, robber- flies (Asilidae), ants, and rodents as predators; and several fungal diseases.
Geographical Distribution
The dearth of collecting records for species of Australian sesiids makes any attempt to analyze distribution patterns both hazardous and tentative.
The absence of life-history data, especially host- plant relationships, adds to the problem, as does the lack of precise knowledge concerning the sesiid fauna of adjacent areas (e.g., New Guinea), which has undoubtedly played a significant role in the development of the Australian sesiid fauna. In spite of these restrictive factors, examination of available data suggests certain probabilities worthy of mention at this point.
Spencer (1896) provided the most generally accepted framework of faunal provinces for the Australian mainland and adjacent Tasmania.
These provinces are illustrated in Map 1 and can be generally characterized both faunistically and ecologically (see Keast et al., 1959; Mackerras, 1970). The Torresian, covering northern Australia, eastern Queensland, and northeastern New South Wales, has a wet to moist, tropical to subtropical climate, patches of rain forest, a general vegeta- tion ranging from wet sclerophyll forest to open grassland, and a fauna that especially reflects
northern elements. The Bassian, extending over southeastern New South Wales, Victoria, south- eastern South Australia, southwestern Australia, and Tasmania, has a moist-temperate climate, veg- etation ranging from southern rain and wet sclerophyll forest to alpine herb fields, and a fauna frequently reflecting a southern element. The Eyrean, consisting of the more arid interior of the continent within the 500 mm isohyet, has a xero- phytic flora and a generally impoverished fauna, frequently reflecting derivation from other prov- inces through progressive adaptation. As is true with any such faunal scheme, it is necessarily arbi- trary, especially at the boundaries, and must be viewed as it relates to the group being studied.
The generalized distribution of Australian sesi- ids in relation to the faunal provinces is illustrated in Map 1. It is immediately apparent that the family occurs primarily in the Torresian area, the only exceptions being Carmenta chrysophanes (Map 6) and Synanthedon tipuliformis (Map 5).
The latter species is a European introduction into Australia and New Zealand and, consequently, does not relate to the present discussion. Thus, within Australia the sesiids display a distinctly tropical and subtropical distribution with virtually no natural penetration of the other faunal prov- inces. Obviously, considering the limited records, this is not a definitive picture; however, it seems unlikely that future studies will significantly alter the overall pattern. In fact, the northern and northwestern portions of Australia have only been minimally sampled for Microlepidoptera thus far, which suggests that the major amount of new infor- mation on sesiids could be expected to come from the Torresian region.
The presently known distribution of sesiids in Australia indicates a definite relationship with the Indo-Malayan area and would appear to be a part of a northern faunal element as summarized by Mackerras (1970). This portion of the Australian fauna includes organisms with ancient ties to the Oriental region, as well as those that have arrived more recently and relate more directly to New Guinea and the Indonesian chain. Common (1970) included the Sesiidae ( = Aegeriidae) among those groups which have reached Australia more re- cently, and our investigations support that deter- mination. The limited penetration of other than the northern tropical and subtropical areas,
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coupled with biological attributes (endophagous boring, hymenopterous mimicry, etc.) that have in other regions allowed radiation into more arid environments, seems a clear indication of recent dispersal into Australia. In the absence of fossil records it is not possible to provide documentation, but it seems to us most likely that the sesiids entered Australia in association with their host- plants during the Indo-Malayan invasion of the late Tertiary. Probable routes of spread during this period are diagrammatically illustrated by
Holloway and Jardine (1968, fig. 21), and the geo- graphical and chronological pattern of this incur- sion has been repeated in numerous publications dealing with many different organisms (Evans, in Keast, et al., 1959). If this view of sesiid disper- sal is valid, it suggests that endemism at the generic level and above should prove minimal and that many, possibly most, of the Australian species will ultimately be found to occur also in New Guinea and adjacent areas, or at least closely related species.
AUSTRALIA
Eyrean Region
NEW ZEALAND
MAP 1.—Distribution of the Family Sesiidae in Australia and New Zealand.
8
Unfortunately, as mentioned at the outset of this discussion, the lack of information concerning the sesiid fauna of New Guinea and islands in the Indonesian chain seriously restricts any judgments concerning extra-Australian relationships. As a result, we are unable to draw more than tentative conclusions concerning endemism at the species level in the Australian sesiids. At the generic level and above, however, no endemism is known to occur. In addition, it is our opinion, based on the examination of available material from the Ori- ental Region, that few, if any, of the species cur- rently known only from Australia are in fact truly endemic. One Australian species, Melittia amboi- nensis, occurs throughout the Indo-Malayan area to India, and while such an extensive range may prove exceptional, it does provide some evidence in support of our position.
As mentioned previously, the only sesiid known to occur in New Zealand is Synanthedon tipuli- formis, introduced along with its hostplant from Europe. First discovered in New Zealand in the late nineteenth century, it now occurs throughout Tasmania and the coastal areas of New South Wales and Victoria in Australia. The absence of other sesiids in New Zealand lends additional support to our contention that the family is a recent arrival in the Australia-New Zealand area.
Classification
During the course of revisionary studies on the Western Hemisphere Sesiidae over the past few years, the authors have attempted to develop a system of higher categories within the family that not only adequately reflects phylogenetic rela- tionships, as we perceive them, within the faunal area of primary concern, but also encompasses the entire family on a world basis. The results of these efforts are, in part, indicated by the system of classification used in this study. In fact, the present study is an outgrowth of a desire to test our higher classification concepts, utilizing a portion of the family far removed from the Western Hemisphere.
The limited number of taxa in the Australian sesiid fauna, coupled with the probability of little or no direct relationship with the Western Hemi- sphere sesiids, seemed to provide a suitable, albeit limited, opportunity for comparison.
In the development of our classification, we have utilized to great benefit three recent works which have also treated this subject in some detail (Niculescu, 1964; MacKay, 1968; Naumann, 1971).
Each of these studies has incorporated new data sources to expand and refine previous concepts, and our efforts have followed a similar pattern.
Although it is not our intention to present a de- tailed account of concepts of higher classification of sesiids in the present paper, a few comments of a general nature seem appropriate.
Without question the most important single work to appear on the family in recent years is that of Naumann (1971). In it Naumann provides a detailed account of previous systems of classifica- tion proposed for the family and compares them at some length with his higher category scheme based on an application of Hennig's principles and methods of phylogenetic systematics. Naumann's classification is based on a detailed study of geni- talia features and wing venation of the type-species of the Holarctic genera and, to a certain degree, of other species in the Palearctic region. In addi- tion, a great deal of nomenclatural clarification and type-species validation is provided for genera worldwide. As a result of these studies, Naumann proposes a monophyletic origin of obscure deriva- tion for the Sesiidae and subdivides the Holarctic sesiid fauna into two subfamilies, Tinthiinae and Sesiinae, recognized phylogenetically as two sister groups. In our studies a broad range of characters including comparative structural morphology of virtually every portion of the adult anatomy, com- parative biological data, and larval morphology, where available, have been utilized for the West- ern Hemisphere sesiids and a significant number of taxa from other parts of the world. As a result of these studies we propose a subdivision of the family into three subfamilies, Tinthiinae, Paran- threninae, and Sesiinae. Obviously, this change in subfamily classification reflects changes at the tri- bal and generic levels as well; however, the overall picture is quite similar to Naumann's arrangement in most instances. In our opinion the classification utilized in this study, due primarily to its basis on a broader range of characters derived from a greater representation of the overall family, pro- vides the most effective arrangement for expressing the relationships of the included taxa.
As mentioned previously, a detailed presenta-
NUMBER 180
tion of the higher categories and phylogeny of the family is intended for inclusion with our studies on the Western Hemisphere sesiids. In the present study all taxa are defined, generally with brief additional comments, in the systematic portion to follow.
One additional observation concerning sesiid classification should be made at this point. Cer- tainly the most surprising aspect encountered in this study was the discovery that all the species of Australian sesiids belong to genera well established in other parts of the world, including the Western Hemisphere. This fact, coupled with data from other studies, tends to reinforce our be- lief that the enormous number of generic names currently applied to sesiids in some areas of the world (e.g., approximately 186 species in 60 genera for the Ethiopian region) bears little or no relation to reality. While the world sesiid fauna is still too poorly known, especially the tropical portion, to make more than tentative observations concerning generic-level endemism, evidence to date suggests that many of the genera are more widely distrib- uted than has been previously recognized. More- over, this situation has no doubt been further aggravated by the tendency of some previous workers to describe newly discovered species in new genera, thereby eliminating the time- consuming necessity of determining whether or not they represent new species in previously described genera.
Taxa Transferred to Other Families During the course of the present study, two taxa were determined to be incorrectly placed in the family Sesiidae. One, a monobasic genus, is trans- ferred to the Stathmopodidae; the other is a species assigned to an appropriate genus in the family Glyphipterygidae.
Family STATHMOPODIDAE
Genus Dolophrosyne Durrant
Dolophrosyne Durrant, 1919:120 [type-species: Dolophrosyne balteata Durrant, 1919].
Dolophrosyne balteata Durant Dolophrosyne balteata Durrant, 1919:121.
TYPE-LOCALITY.—Queensland, Australia.
TYPE.—Holotype male, British Museum (Natu- ral History).
DISCUSSION.—This genus and its single included species is here transferred to the family Stathmo- podidae. The type-specimen was studied and proved not to be a sesiid. Although precise place- ment must await careful study of the family Stath- mopodidae, the species appears nearest the genera Snellenia and Pseudaegeria.
Family GLYPHIPTERYGIDAE Genus Sagalassa Walker
Sagalassa Walker, 1856:8 [type-species: Sagalassa robusta Walker, 1856].
Sagalassa homotona (Swinhoe), new combination Balataea homotona Swinhoe, 1892:36.
TYPE-LOCALITY.—Australia.
TYPE.—Holotype male, University Museum, Hope Department of Entomology, Oxford, England.
DISCUSSION.—This species is here transferred to the genus Sagalassa in the family Glyphipterygidae.
Examination of the type-specimen clearly indicates that the species was improperly placed in the Sesiidae. Turner (1922:59) indicated in a foot- note to his tabulation of Australian sesiids that this species belonged in the genus Miscera Walker ( = Sagalassa Walker); however, this transfer apparently went unnoticed and the species has continued to be included in the family until the present time.
Checklist Family Sesiidae
Subfamily Tinthiinae Tribe Pennisetiini
Pennisetia Dehne
= Anthrenoptera Swinhoe
= Lophocnema Turner
= Diapyra Turner
P. igniflua (Lucas) P. eusphyra (Turner) Tribe Tinthiini
Tinthia Walker
T. xanthospila Hampson Subfamily Paranthreninae
Tribe Paranthrenini Albuna Edwards
= Harmonic Edwards
= Parharmonia Beutenmuller A. carulifera (Hampson)
= coracodes Turner A. isozona (Meyrick) A. oberthuri (LeCerf)
= terrible Turner A. zoniota (Turner) Subfamily Sesiinae
Tribe Melittiini Melittia Hubner
= Poderis Boisduval Af. atnboinensis Felder
= doddi LeCerf
= thaumasia Turner Af. chalybescens Miskin
= proserpina Hampson Tribe Synanthedonini
Synanthedon Hubner S. tipuliformis (Clerck) 5. cupreifascia (Miskin) Carmen ta Edwards
C. chrysophanes (Meyrick)
= melanocera Hampson
= panyasis Druce
= caieta Druce
C. commoni Duckworth and Eichlin C. xanthogyna Hampson
Family SESHDAE
SESIIDAE Boisduval, 1828:29 [Sesiariae; type-genus: Sesia Fabricius, 1775].
AECERIIDAE Stephens, 1829:34.—Naumann, 1971:9 [synonymy].
[Refer to Dalle Torre and Strand, 1925:1-3, for detailed bibliography.]
DIAGNOSTIC CHARACTERS.—Small to fairly large moths, forewing length 5-30 mm. Head: Eyes rela- tively large compared to width and length of head;
naked, ringed with short yellow or white scales.
Ocelli present. Pilifers with rather strong, spine- like setae of varying length. Mandibles small, flat- tened, and somewhat hatchet-shaped. Proboscis most often elongate and functional but may be rudimentary and nonfunctional in some species.
Maxillary palpus small, 1- to 3-segmented, most
species with two segments. Labial palpus 3- segmented, upcurved. Antenna of most species clavate, tapered to point apically, terminated by small scale tuft; males of all but few species with setae ventrally, pectinate in some groups. Thorax:
Dorsoanterior region of preepimeron with hollow, weakly sclerotized, baglike protuberance on all species except those in the Tinthiinae. Abdomen:
Generally elongate, slender, tapering posteriorly, often narrowed at base with anal scale tuft most developed on males. Legs: Generally elongate, slender; first tarsal segment of hindleg elongate, about one-half length of tibia; tibia with scale tufts near spurs. Forewing: Elongate, narrow, varying from mostly hyaline to entirely opaque.
Veins: R generally with five branches, R4 and R-, most often stalked; M lost in cell, often with three branches; Cu with two branches. Dorsal mar- gin of wing folded down, interlocking with up- folded costal margin of hindwing (see discussion of wing-locking mechanism). Hindwing: Elongate narrow, somewhat broader than forewing, varying from hyaline to opaque depending on the species.
Frenulum single in both sexes. R veins combined (Rs), often hidden in fold; M veins separate; M3 and Cuj joined before or after crossvein. Anal Veins: 1A usually degenerate, present in primitive species; 2A present; 3A either short, fused to base of 2A or absent; 4A most often present, absent in primitive species. Male Genitalia: Uncus simple in few groups, more often modified and/or fused with tegumen, with lateral setaceous pads or with elongate, paired socii clothed with specialized bi- furcate setae; gnathos absent or present in various forms; tegumen simple, reduced, or modified and fused with uncus; vinculum somewhat ring-shaped, extended anteriorly as saccus of varying length and thickness; valva varies from small and quad- rangular to elongate and slender, with either sim- ple setae or combination of simple setae of varying shapes and sizes and arrangements of bifurcate or multi-furcate setae; specialized saccular ridge pres- ent or absent. Aedeagus generally elongate, very slender, slightly bulbous at base. Female Genitalia:
Papillae anales generally small and narrow; ostium bursae situated ventrally between sclerites of ab- dominal segment 8, or in intersegmental membrane between 7 and 8, or on posterior margin of abdomi- nal segment 7; ductus bursae most often elongate, narrow, with sclerotization for the most part con-
NUMBER 180 1 1
Key to the Species and Higher Categories
1. Antennae filiform, without apical scale tuft (Tinthiinae) 2 Antennae clavate, with apical scale tuft 4 2. Hindwing with vein M3 arising from Cux well basad of crossvein (Tinthiini)
Tinthia xanthospila Hindwing with veins M3 and CUj long-stalked distad of crossvein (Pennisetiini: Pennisetia) 3 3. Forewing opaque, dorsally with at least some orange Pennisetia igniflua Forewing with hyaline area apically, dorsally with some straw-yellow and tan, without orange Pennisetia eusphyra 4. Hindwing with vein M3 arising from Cux basad of crossvein; vein 3A absent; forewing with stalk of vein R4 plus R5 more than one-half total length of R< or R5 (Paranthreninae:
Paranthrenini: Albuna) 5 Hindwing with vein M3 arising from Cut distad of crossvein, or if basad of crossvein then vein 3A present; forewing with stalk of vein R4 plus R5 not more than one-half total
length of R4 or R5 (Sesiinae) 8
5. Forewing mostly or entirely opaque with at least some blue iridescence dorsally
Albuna carulifera Forewing mostly hyaline or if mostly opaque, then orange, without blue iridescence 6 6. Abdomen dorsally with one narrow white band; head with front white with brown-black;
labial palpus white with brown-black Albuna zoniota Abdomen dorsally with two or more yellow or yellow-orange bands; head with front yellow;
labial palpus yellow or orange with brown-black 7 7. Head with vertex roughened posteriorly, labial palpus strongly roughened laterally (males) or ventrally (females); thorax with collar yellow; female with cell opaque on forewing, discal spot orange or not defined; abdomen dorsally with yellow narrowly banded on segment 4 and mostly yellow on 6, ventrally without yellow Albuna oberthuri Head with vertex smooth, labial palpus mostly smooth; thorax with collar brown-black with some yellow-orange laterally; female with cell hyaline on forewing, discal spot brown-black; abdomen dorsally with broad yellow-orange bands on segments 2, 4, and 6, ventrally with all segments broadly banded yellow-orange Albuna isozona 8. Hindwing with vein M3 arising from Cux well basad of crossvein; scale-plate present beneath scape of antenna; hindlegs very strongly tufted (Melittiini: Melittia) 9 Hindwing with vein Ms arising from Cux distad of crossvein; scale-plate beneath scape absent; hindlegs untufted or with slight tufting confined primarily to region of tibial spurs (Synanthedonini: Carmenta and Synanthedon) 10 9. Forewing at least one-half opaque, with broad apical margin and broad discal spot; abdomen
white or solid brown-black ventrally; tufts of hindlegs without orange
Melittia chalybescens Forewing mostly hyaline, with narrow apical margin and discal spot; abdomen yellow or
yellow and orange ventrally; tufts of hindlegs strongly orange laterally
Melittia amboinensis
10. Forewing with vein R2 missing Carmenta commoni
Forewing with vein R2 present, often very close to R,, occasionally fused apically 11 11. Head with antennae mostly orange, front mostly white, labial palpus flattened ventrally, orange with white; forewing with discal spot mostly orange Synanthedon cupreifascia Head with antennae not orange, front brown-black with some white laterally, labial palpus not flattened ventrally, yellow with brown-black; forewing with discal spot brown-black .12 12. Abdomen brown-black ventrally, occasionally some yellow on posterior margin of segment 4 Synanthedon tipuliformis Abdomen mostly or entirely yellow ventrally 13 13. Forewing mostly hyaline between veins R» and Rs Carmenta xanthogyna Forewing opaque between veins R4 and R3 Carmenta chrysophanes
12
fined to posterior portion; ductus seminalis origi- nates from various points along ductus bursae, most often toward posterior end; corpus bursae generally small, obovate and membranous with convolu- tions or signa in some species. Eggs: Generally pale brown to chestnut brown, ovate, disc-shaped, flat, or slightly concave ventrally; surface weakly sculp- tured with minute shagreening in hexagonal de- signs. Larvae: Pale, color restricted to head and prothoracic shield; ocelli I to IV arranged in trapezoid and remote from V and VI; crochets uniordinal in two transverse rows. Pupae: Dorsal abdominal spines in double rows on segments 2 to 7 on male and 2 to 6 on female, single row on 7 in female, both sexes with single rows on 8 to 10;
large spines ventrad on 10; cremaster absent; ab- dominal segments freely movable; head sometimes modified with cutting device, somewhat chisel-like.
Subfamily TINTHHNAE LeCerf
TINTHIINAE LeCerf, 1917:48 [type-genus: Tinthia Walker, 1864].
BEMBECIINAE Niculescu, 1964:42 Naumann, 1971:47 [syn- onymy].
ZENODOXINAE MacKay, 1968:5.—Naumann, 1971:47 [synon- ymy].
DIAGNOSTIC CHARACTERS.—Head with maxillary palpus (Figure If) 1-segmented; antennae filiform (Figure \b), never clavate, without terminal scale tuft. Forewing (Figures 2 and 3) with veins R4 and Rr, stalked or unstalked; Cu2 absent or very short. Hindwing with 1A nearly or entirely de- veloped, 2A present, 3A and 4A absent. Male genitalia with valva having only simple setae, with- out saccular ridges; anellus well developed, sleeve- like; base of saccus and vinculum broad, thick;
tegumen simple; uncus a simple projection or re- duced to pair of setose knobs; gnathos absent.
DISCUSSION.—This subfamily is represented in the Australian fauna by three species in two genera of two tribes, Pennisetiini and Tinthiini. The Tin- thiinae, in our opinion, is the least specialized sub- family of the Sesiidae based on the retention of certain structures, such as 1A in the hindwing, and the absence of many specializations and modifica- tions which occur in the other two subfamilies.
The subfamily appears to be best represented, on a world basis, in eastern Asia.
Tribe PENNISETIINI Naumann
PENNISETIINI Naumann, 1971:55 [new name for Bembeciini Niculescu; type-genus: Pennisetia Dehne, 1850].
DIAGNOSTIC CHARACTERS.—Head with maxillary palpus 1-segmented; antennae of male with long cilia ventrally (strongly bipectinate on species in other faunal regions). Forewing with stalk of veins R4 and R5 less than one-half total length of either vein; Cu2 absent. Hindwing with veins M3 and Cui long-stalked distad of crossvein.
DISCUSSION.—This tribe is represented in the Australia-New Zealand area by two species in the genus Pennisetia.
Genus Pennisetia Dehne
Pennisetia Dehne, 1850:28 [type-species: Pennisetia anomala Dehne, 1850, a synonym of Sesia hylaeiformis Laspeyres, 1801].
Anthrenoptera Swinhoe, 1892:35 [type-species: Sphecia con- tracta Walker, 1856] Naumann, 1971:56 [synonymy].
Lophocnema Turner, 1917:78 [type-species: Lophocnema eusphyra Turner, 1917]. [New synonymy.]
Diapyra Turner, 1917:79 [type-species: Sesia igniflua Lucas, 1894]. [New synonymy.]
Pennisetia igniflua (Lucas), new combination
FIGURES \b, If, 2, 9, 19, 29, 30; MAP 2
Sesia igniflua Lucas, 1894:133.
Diapyra igniflua (Lucas) Turner, 1917:79; 1922:62.
Glossecia igniflua (Lucas) Hampson, 1919:113.
MALE.—Head with vertex brown-black; occipital fringe brown-black, white laterally; antennae brown-black; front gray-black; labial palpus dorso- ventrally flattened, white, third segment brown- black. Thorax mostly brown-black, with narrow, subdorsal, orange stripes on posterior one-half of mesothorax; strongly white beneath wings; lateral tufts of metathorax white. Abdomen somewhat dorsoventrally flattened, widest at segment 4 and tapered to blunt point posteriorly; dorsally brown- black with white laterally on segment 4, and with median tuft on segment 3; ventrally mostly white;
anal tuft undeveloped. Legs with coxae and femora mostly white; tibiae laterally mostly brown-black, strongly spined medially and distally; hindtibia with basal one-half white, spurs mostly white;
NUMBER 180 13 tarsi mostly brown-black with white basally on
most segments, and strongly spined distally on first segment. Forewing opaque; brown-black, strongly orange on basal one-half, and orange forming large spot in apical one-third; ventrally without orange; fringe tipped with white. Hind- wing mostly hyaline; margin variously suffused, brown-black; no discal spot; brown-black scales on veins raised, strongest on Cu veins; ventrally with some white powdering apically; fringe brown- black with white at base of scales. Wing length of
male, 6 mm. Male genitalia (Figure 9) with valva simple, somewhat truncate apically, produced and expanded basally, clothed on apical one-half with simple setae; saccus elongate; anellus elongate, tubular, somewhat sclerotized ventrally; uncus broad, terminating as two lateral setose knobs;
subscaphium broad, platelike; aedeagus elongate, slender, only slightly thicker at base, as long as valva and saccus combined.
FEMALE.—Differs from the male by the following:
mesothorax solid orange dorsomedially; abdomen
1. Penm'setia eusphyra 2. P. Igniflua 3. Tinthia xanthospila
NEW ZEALAND
MAP 2
14
dorsally with segment 4 mostly white except for brown-black medially, narrowly white on anterior margin of segment 3, often also on segment 2, some white variously on other segments; forewing dor- sally with basal two-thirds mostly orange, apical orange spot more extensive than on male, white on tornus; wing length of female, 6-7 mm. Female genitalia (Figure 19) with ostium bursae a small circular opening ventromedially on posterior edge of segment 7; initial short section of ductus bursae slightly sclerotized to origin of ductus seminalis, then slightly expanded, continuing as membra- nous, elongate tube to obovate corpus bursae; sig- num of corpus bursae ribbon-like, about two-thirds length of corpus bursae, well sclerotized and pigmented.
TYPE.—The following two syn types from the South Australian Museum were examined: (1)
"B"; "S. A. Museum Specimens" (female). (2)
"B"; "TYPE $"; "S. A. Museum Specimens,"
(female). Lucas (1894) in the original descrip- tion of this species wrote: " $ $ 14-15 mm."; "Bris- bane; in scrub."; "Dr. Turner succeeded in captur- ing seven specimens this year and has courteously given me a pair." Accompanying above-listed syntypes is a single label, "Diapyra igniflua Lucas, 114869, Brisbane, Type." Lucas apparently thought the smaller of the two specimens, syntype I, was the male of the pair to which he referred, when in fact both specimens are females. Without additional information to the contrary, it is our opinion that the above-listed syntypes represent the original type-series. The better of the two specimens, the first female syntype listed above from the South Australian Museum, has been se- lected, labeled, and is presently designated the lectotype.
TYPE-LOCALITY.—Brisbane, Queensland, Austra- lia.
HOST PLANT.—Elaeocarpus grandis (Elaeocar- paceae).
DISCUSSION.—All 10 specimens examined were from Brisbane. Both Lucas (1894) and Turner (1917) indicated the host of this species as Elaeo- carpus grandis; however, Tillyard (1926) reported the host-plant to be Santalum acuminatum. We have found no record in the literature which verifies this statement and assume the confusion stems from the fact that the two species of plants are both commonly called "quandong trees." The
common name was mentioned by Lucas along with the scientific name in the initial publication, and since both Lucas and Turner indicated that the species they were rearing from was Elaeocarpus grandis, we consider Tillyard's record in error.
Although P. igniflua is only known from Bris- bane, the distribution of its known host-plant suggests that it will ultimately be found to occur in rain-forest areas in Queensland and the North- ern Territory.
Pennisetia eusphyra (Turner), new combination
FIGURES 8, 28; MAP 2
Lophocnema eusphyra Turner, 1917:79.
MALE.—Head with vertex brown-black, some straw-yellow laterally; occipital fringe brown-black, white laterally; antennae brown-black, strongly powdered tan dorsally; front brown-black, tan laterally and ventrally; labial palpus dorsoventrally flattened, tan with white ventrad. Thorax brown- black with straw-yellow surrounding wing base, extending dorsolaterally to metathorax; ventrally brown-black with tan mixed; metathorax with lateral tufts brown-black. Abdomen dorsally brown- black with segments 4 and 7 golden-yellow, seg- ment 1 white anteriorly, medial tufts on segments 3 to 6 tan except for straw-yellow on 4; ventrally with white on segments 2 to 4; anal tuft golden- yellow dorsally, tan ventrally. Legs tan, with much brown-black on coxae and femora; foretibia strongly roughened; dorsally midtibia strongly tufted basad and distad with paler, spinelike scales; hindtibia very strongly tufted from middle spurs to apex of tibia dorsomedially, broadly fan- shaped at apical spurs; first tarsal segment tufted distad. Forewing opaque except for hyaline area distad of discal spot, extending from vein R4+r, to nearly M3; brown-black mixed with straw- yellow and tan, plus few pale blue scales medially, and some orange apically; forewing ventrally mostly brown-black. Hindwing hyaline with very narrow, brown-black margins; scales on veins raised as on P. igniflua, brown-black; no discal spot. Wing length of male, 6 mm. Male genitalia (Figure 8) similar to P. igniflua, but with aedea- gus longer than combined length of saccus and valva; saccus shorter, and valva more rounded apically than for P. igniflua.
NUMBER 180 15
FEMALE.—Generally larger, with more pale orange on forewing apically, tibiae not as strongly tufted, otherwise the same as for the male. Wing length of female, 8-9 mm. Female genitalia much the same as P. igniflua, but apparently lacking the signum on the corpus bursae.
TYPE.—The following two syntypes from the Australian National Insect Collection were exam- ined: (1) "Lophocnema eusphyra Turn., TYPE";
"Cairns dist., F. P. Dodd"; "LECTOTYPE. Lop- hocnema eusphyra Turner, 1917, designated by";
"Aust, Nat. Ins. Coll."; "Genitalia Slide By T. D.
Eichlin, USNM 76095 $" (male). (2) "Cairns dist., F. P. Dodd"; "PARALECTOTYPE, Lophoc- nema eusphyra Turner, 1917, designated by";
"Aust. Nat. Ins. Coll."; "Genitalia Slide By T. D.
Eichlin, USNM 76096 9 " (female). In the origi- nal description Turner (1917) wrote, "N. Q. Ku- randa near Cairns; two specimens received from Mr. F. P. Dodd." He described both sexes. Another pair of specimens examined from the National Museum of Victoria, Melbourne, was labeled,
"Cairns dist., F. P. Dodd", but it is our opinion that the first pair listed above is the actual syn- typic series indicated by Turner. The male syn- type listed above from the Australian National Insect Collection has been selected, labeled, and is presently designated as the lectotype.
TYPE-LOCALITY.—Kuranda, Queensland, Austra- lia, near Cairns.
HOST PLANT.—Unknown.
DISCUSSION.—Only four specimens of this species were available for study. Morphologically this species is very similar to P. igniflua but may be easily distinguished by differences in the macula- tion of the wings, legs, and abdomen, which have been detailed above. Turner (1917) placed the two species in separate genera based primarily on the erroneous observation that the males of P.
eusphyra lack a proboscis. In fact, though tightly coiled on some specimens, both sexes of each species have a well-developed proboscis. Nothing is known of the biology of P. eusphyra; however, considering the similarities with P. igniflua, it seems likely that the species will ultimately be found on one or more of the numerous species of Elaeocarpus found throughout the rain-forest areas from northern New South Wales north along the coast of Queensland.
Tribe TINTHHNI LeCerf
TINTHIINI LeCerf, 1917:148 [type-genus: Tinthia Walker, 1864].
ZENODOXINI MacKay, 1968:5.—Naumann, 1971:49 [synonymy].
DIAGNOSTIC CHARACTERS.—Head with maxillary palpus 1-segmented; antennae of male ciliate ven- trally; eyes comparatively smaller than other species of Sesiidae. Forewing with veins R4 and R5 short-stalked (unstalked on most species else- where); Cu2 absent or very short. Hindwing with vein M3 arising from Cux well basad of crossvein.
DISCUSSION.—Only one species in this tribe, a member of the genus Tinthia, is known to occur in the Australian and New Zealand fauna.
Genus Tinthia Walker
Tinthia Walker, 1864:23 [type-species: Tinthia varipes Walker, 1864].
Tinthia xanthospila Hampson
FIGURE 3; M A P 2
Tinthia xanthospila Hampson, 1919:115.
Tinthia xanthopila [sic] Hampson, Dalla Torre, and Strand, 1925:183.
MALE.—Head with vertex orange posteriorly, brown-black anteriorly; occipital fringe orange;
antennae brown-black (broken); front brown- black; labial palpus smooth, dorsoventrally flat- tened, orange. Thorax brown-black; collar orange, tegulae mostly orange; metathorax orange. Abdo- men missing. Legs with foreleg orange except dorsally on tibia; midleg mostly orange with brown-black dorsally on femur, dorsally and dis- tally on tibia, and all of first tarsal segment; hind- leg like midleg but tarsal segments mostly brown- black. Forewing opaque, brown-black with orange spot centered on distal one-half; ventrally orange, also strongly powdered basally. Hindwing hyaline, brown-black, margins narrow though slightly dif- fuse. Wing length of male, 7 mm. Male genitalia unknown.
FEMALE.—Unknown.
TYPE.—Holotype: Male in the British Museum (Natural History): "Cedar Bay, Queensland, Meek, 1894, Rthschld. 17775"; "Tinthia xanthos- pila Hmpsn., type $ "; "Type" (abdomen missing).
16
TYPE-LOCALITY.—Cedar Bay, Queensland, Aus-
tralia.
HOST-PLANT.—Unknown.
DISCUSSION.—This species is known only from the type-specimen, which is in imperfect condition.
A more detailed characterization of the species must await the acquisition of additional speci- mens and, hopefully, data concerning its life history.
Subfamily PARANTHRENINAE Niculescu
PARANTHRENINAE Niculescu, 1964:38 [type-genus: Paranthrene Hubner, 1819].
SESIINAE Boisduval, Naumann, 1971:58 [in part].
DIAGNOSTIC CHARACTERS.—Head with maxillary palpus (Figure Id) 2-segmented, second segment large, often indented in middle; antennae (Figure la) clavate with terminal tuft of scales, ciliate ventrally on male (often bipectinate on species of other faunal areas); eyes proportionately larger than for species of Tinthiinae. Forewing (Figure 4) with veins R4 and R5 long-stalked, stalk longer than one-half total length of R4 or R5; Cu2 pres- ent, often as long as Cux. Hindwing with vein M3 arising from Cuj just basad of crossvein; 1A degenerate (to observe, scales must be removed), 2A and 3A coincident except at base, 4A present.
Male genitalia with valva having scales multifur- cate dorsally and setaceous ventrally and apically, with median area unsealed or with thick, dark scales in saccular region; tegumen reduced;
gnathos small, rounded or bifurcate, often with teethlike processes on ventral margin; uncus wide, elongate, about three to five times longer than tegumen, clothed lateroapically with long seta- ceous scales, mostly bilobed apically; vinculum narrow, saccus relatively short.
DISCUSSION.—Included in this subfamily for Australia-New Zealand are four species of Albuna in the tribe Paranthrenini. The subfamily Paran- threninae occurs worldwide with its greatest diver- sity appearing in the Eastern Hemisphere.
Tribe PARANTHRENINI Niculescu
PARANTHRENINI Niculescu, 1964:38 [type-genus: Paranthrene Hubner, 1819] .
DIAGNOSTIC CHARACTERS.—Refer to description
given for subfamily. The data provided for the subfamily will suffice for distinguishing the nomi- nal tribe since no other tribes are represented in the fauna under consideration.
DISCUSSION.—This tribe in the area under study contains one genus, Albuna, with four included species.
Genus Albuna Edwards
Albuna Edwards, 1881:186 [type-species: Aegeria hylotomi- f or mis Walker, 1856, now considered a synonym of Aegeria pyramidalis Walker, 1856].
Harmonia Edwards, 1882:54 [type-species: Harmonia mor- risoni Edwards, 1882, now considered a synonym of Car- menta fraxini Edwards, 1881].
Parharmonia Beutenmuller, 1894:89 [new name for Harmonia Edwards].
DISCUSSION.—This genus is closely related to Paranthrene, but may be differentiated by the fol- lowing characters: antennae of male ciliate ven- trally but not pectinate; forewing with bases of vein R3 and stalk of veins R4 and R5 connected by a crossvein, not connate; male genitalia with gnathos having spinelike processes on ventral mar- gin, and with lateral portions of uncus naked or sparsely clothed with setaceous scales (Australian species).
Albuna carulifera (Hampson), new combination
FIGURES 10, 20, 31, 32; MAP 3
Paranthrene carulifera Hampson, 1919:108.
Paranthrene caerulifera Hampson Turner, 1922:62 [invalid emendation].
Trochilium coracodes Turner, 1922:61. [New synonymy.]
Conopia coracodes (Turner).—Dalla Torre, and Strand, 1925:
109.
MALE.—Head with vertex brown-black; occipital fringe brown-black dorsally, white laterally; anten- nae brown-black, scape white; front brown-black with white laterally; labial palpus broadly roughened, brown-black, strongly mixed with white dorsally and ventrally. Thorax brown-black, mixed with white beneath wings and laterally above wings. Abdomen brown-black, lightly pow- dered pale yellow and white, scattered laterally and ventrally on each segment; anal tuft narrowly edged with white laterally. Legs brown-black;
